ONTOGENY OF THE STEM-NODE-LEAF VASCULAR CONTINUUM OF AUSTROBAILEYA

Developmental study of the stem-node-leaf vascular continuum of Austrobaileya scandens White reveals that the vasculature within each leaf originates from a single procambial strand, that becomes separated into two strands only at the junction of leaf and stem. At lower levels in the stem the two strands become incorporated into independent portions of the stele. At later stages of development the solitary vascular bundle within the young leaf undergoes considerable lateral growth, resulting in an essentially continuous arc of vascular tissue. Ontogenetic evidence indicates that the vascular bundle in the midrib of the lamina should be regarded as a fundamentally single bundle and not interpreted as two bundles that have undergone various degrees of secondary fusion. A condition of two totally separate bundles extending the entire length of the leaf was not encountered. Our observations confirm the characterization of Austrobaileya as an example of “second rank” level of leaf vasculature. Nodal anatomy emphasizes the extremely isolated taxonomic position of Austrobaileya within the primitive dicotyledons.     

MAJOR CLADES OF THE ANGIOSPERMS

Abstract— Our knowledge of fundamental angiosperm interrelationships is still very incomplete. The absence of a narrowly circumscribed gymnosperm outgroup, ideally the sister group, makes character evaluation, necessary for a cladistic analysis, difficult. According to current views the superorder Magnoliiflorae with a number of other groups, for example the monocotyledons, may represent a complex of families near the base of the angiosperms. Interrelationships of groups within the monocotyledons are much better understood than those between groups within the dicotyledons. A cladogram of monocotyledon orders based on earlier work by R. Dahlgren, H. T. Clifford, and F. N. Rasmussen is presented. A data matrix for a sample of the angiosperms with 61 characters for 49 taxa, mostly magnoliifloran and related families, is presented. The characters are polarized mainly according to the current view that the primitive angiosperm morphotype is a woody dicotyledon with strobiloid flowers. As an alternative the matrix is adjusted following W. C. Burger's conjecture that the primitive angiosperm was a herbaceous monocotyledon with trimerous flowers. Both matrices were run in a computerized parsimony analysis, resulting in numerous equally parsimonious solutions. This result is illustrative of the great homoplasy in the available character information, and also of how little actually is known about fundamental angiosperm interrelationships or phylogeny.     

SHOOT VASCULAR SYSTEM AND PHYLLOTAXIS OF CASUARINA (CASUARINACEAE)

In species of Casuarina with multileaved whorls, each stem vascular bundle divides radially into two at the site of a leaf trace separation, and the same two bundles rejoin acropetally to where the trace supplies a leaf. Such divisions are divisions of a single vascular bundle, and the rejoining of bundles forms a single bundle. Proposals that the extant primary vascular systems of dicotyledons may have been derived as in conifers are incorrect in so far as Casuarina is concerned, or the system has evolved beyond that so far proposed for dicotyledons. Reasons are offered, however, for considering that fernlike leaf gaps are not present. Leaf traces supply leaves at the first nodes distal to their origins. The ways by which an increase or decrease of stem bundles occur are described. Phyllotactic patterns range from helical (rare) to whorled. In the embryo, where leaves occur decussately, of certain species with multileaved whorls, and in the shoot apices of species with tetramerous whorls, slight differences in the levels of leaf attachments and the bending of leaf traces indicate the probable evolution of extant whorled phyllotaxies from one or more helical arrangements. Stages in the evolution are suggested. The leaves in most species with multileaved whorls are in true whorls. The original periderm of branchlets lies internally to the internodal traces and chlorenchyma, but is otherwise external to the vascular system. It is concluded that each leaf originates at its level of separation from the axis despite several structural features suggesting that the leaf bases have become congenitally adnate to the stem.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE SENSU LATO. XVII. FLORAL ANATOMY OF ONDINEA PURPUREA SUBSPECIES PURPUREA (NYMPHAEACEAE)

Classification and phylogeny of the Nymphaeaceae are unresolved. This study provides floral anatomical data that will assist in elucidating generic interrelationships and systematic relationships to other taxa of angiosperms. The floral anatomy of Ondinea purpurea den Hartog subsp. purpurea has been examined utilizing light microscopy. The peduncle possesses stelar vascular bundle complexes and cortical vascular bundles. Cortical bundles terminate within the peduncle. Each bundle complex consists of 2 collateral bundles on the same radius, the inner bundle inverted; 2 protoxylary lacunae occur yet differ in structure and function. Progressing acropetally, the inner xylary lacunae become discrete mesarch strands surrounded centrifugally by a vascular cylinder formed by divisions and anastomosing of the bundle complexes. Together these become the massive receptacular vascular plexus. The plexus provides collateral traces to the floral organs. Each sepal receives 3 traces that separate from the plexus as 1–3 lateral traces. Petals are absent and no vestigial petal traces have been observed. Distally, the plexus forms several large strands of connate gynoecial and androecial traces termed the principal vascular bundles (PVBs). Ventral veins separate from the PVBs and the latter extend acropetally through the outer ovary wall. Branches of the ventrals and PVBs contribute to septal vascular reticula from which each ovule is supplied by one vascular bundle. Each stamen receives 1 trace from branches of the PVBs. The ventrals and PVBs terminate within the carpellary lobes. A comparative anatomical study is offered that supports the inclusion of Ondinea in the Nymphaeaceae sensu stricto.     

HISTOLOGICAL STUDIES ON THE COLEOPTILE. I. TISSUE AND CELL TYPES IN THE COLEOPTILE TIP

The uppermost 1-4 mm of 25-mm coleoptiles of oats and wheat have been studied at the optical microscope level, using newer histological methods and sections 1-4 μ thick. The outer epidermal wall, which shows very fine wrinkling, is continuous with the thinner wall of the inner epidermis through the pore. The cells of both epidermal layers have acidophilic cytoplasm with long transvacuolar strands. Both inner and outer epidermis have stomata, those of the outer epidermis having kidney-shaped guard cells like those of dicotyledons. The guard-cell walls are lignified in their inner layers only and are thinly cuticularized. In the vascular bundles the sieve tubes terminate apically about 250 μ below the end of the xylem; the xylem in turn terminates about 400 μ below the extreme apex. A number of clearly undifferentiated cells, with highly basophilic cytoplasm and many mitochondria, separate the xylem elements from the inner epidermis. Towards the outer epidermis there are a few sieve elements, each of which is associated with a special cell having an elongated nucleus supported on fine cytoplasmic strands. The parenchyma of both the tip and the shaft of the coleoptile are generally interpenetrated by air-spaces, but where they are adjacent to the inner epidermis there is heavy interposition of readily stained intercellular material, especially in Triticum. Plastids are widely distributed throughout the tissue, but their greening in light takes place preferentially towards the phloem side of the vascular bundles. The observations are discussed in reference to earlier literature and with regard to the function of the coleoptile as a protecting and guiding organ for the shoot system of the seedling.     

VASCULAR CONSTRUCTION AND DEVELOPMENT IN THE AERIAL STEM OF PRIONIUM (JUNCACEAE)

The aerial stem of Prionium has been studied by motion-picture analysis which permits the reliable tracing of one among hundreds of vascular strands throughout long series of transverse sections. By plotting the path of many bundles in the mature stem, a quantitative, 3-dimensional analysis of their distribution has been made, and by repeating this in the apical region an understanding of vascular development has been achieved. In the mature stem axial continuity is maintained by a vertical bundle which branches from each leaf trace just before this enters the leaf base. Lateral continuity results from bridges which link leaf traces with nearby vertical bundles. Development of the provascular system involves a meristematic cap into which the blind ends of vertical bundles can be followed. Leaf traces are produced continuously in association with developing leaf primordia for a period of over 30 plastochrones; they connect with the vertical bundles in the meristematic cap and so establish the essential vascular configuration which is later reorientated through about 90° by overall growth of the crown. The last bundles to differentiate from the leaf do so outside the meristematic cap and thus fail to make contact with the axial system; they appear in the mature axis as blind-ending cortical bundles. Prionium is only distantly related to palms and its vascular histology is quite different. Nevertheless, the course of vascular bundles and the origin of this pattern in the stem resembles that of a palm. It is suggested that we are examining the fundamental pattern of vascular development in large monocotyledons.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. I. NYMPHAEA SUBGENERA CASTALIA AND HYDROCALLIS

The vascular system in the stems of Nymphaea odorata and N. mexicana subgenus Castalia, and N. blanda subgenus Hydrocallis consists of continuing axial stem bundles with eight being the usual number. The stem bundles are concentric and xylem maturation is mesarch. Xylem elements consist of tracheids with spirally or weakly reticulated secondary wall thickenings. The phloem is made up of companion cells and short sieve tube members with simple sieve plates that are nearly transverse. At the node each leaf is supplied with two lateral leaf traces and a median leaf trace. A root trace is also present and supplies a series of adventitious roots borne on the leaf base. Flowers and vegetative buds develop directly from the apical meristem and occupy leaf sites in a single genetic spiral. Each flower or vegetative bud is related to a leaf through specific spatial and vascular association. The related leaf is separated from the related flower by three members of the genetic spiral and occupies an adjacent orthostichy. Vascular tissue for the related flower arises from the inner surfaces of the four stem bundles supplying leaf traces to the related leaf and extends through the pith to the flower or vegetative bud via a peduncle fusion bundle. The vascular system organization in the investigated species of Castalia and Hydrocallis is not typically monocotyledonous or dicotyledonous, nor can it be considered transitional between them. The ontogeny of the vascular system is similar to typical dicotyledons and the investigated species of Nymphaea can, therefore, be considered to represent highly specialized and modified dicotyledons.     

VASCULAR ORGANIZATION IN SHOOTS OF CACTACEAE. I. DEVELOPMENT AND MORPHOLOGY OF PRIMARY VASCULATURE IN PERESKIOIDEAE AND OPUNTIOIDEAE

Primary shoot vasculature has been studied for 31 species of Pereskioideae and Opuntioideae from serial transections and stained, decorticated shoot tips. The eustele of all species is interpreted as consisting of sympodia, one for each orthostichy. A sympodium is composed of a vertically continuous axial bundle from which arise leaf- and areole-trace bundles and, in many species, accessory bundles and bridges between axial bundles. Provascular strands for leaf traces and axial bundles are initiated acropetally and continuously within the residual meristem, but differentiation of procambium for areole traces and bridges is delayed until primordia form on axillary buds. The differentiation patterns of primary phloem and xylem are those typically found in other dicotyledons. In all species vascular supply for a leaf is principally derived from only one procambial bundle that arises from axial bundles, whereas traces from two axial bundles supply the axillary bud. Two structural patterns of primary vasculature are found in the species examined. In four species of Pereskia that possess the least specialized wood in the stem, primary vascular systems are open, and leaf traces are mostly multipartite, arising from one axial bundle. In other Pereskioideae and Opuntioideae the vascular systems are closed through a bridge at each node that arises near the base of each leaf, and leaf traces are generally bipartite or single. Vascular systems in Pereskiopsis are relatively simple as compared to the complex vasculature of Opuntia, in which a vascular network is formed at each node by fusion of two sympodia and a leaf trace with areole traces and numerous accessory bundles. Variations in nodal structure correlate well with differences in external shoot morphology. Previous reports that cacti have typical 2-trace, unilacunar nodal structure are probably incorrect. Pereskioideae and Opuntioideae have no additional medullary or cortical systems.     

VASCULAR PATTERNS IN STEMS OF THE CYCLANTHACEAE

A survey was made of the distribution of stem vascular bundles in representatives of ten genera of the tropical monocotyledonous family Cyclanthaceae. Films of series of serial transverse sections were used to reconstruct the stem vasculature. Each leaf trace, followed in a basipetal direction from its level of insertion at the stem periphery, describes an obliquely downward course, initially contacting from 1 to 4 (or more) existing axial bundles. The associated bundles form a compound vascular bundle in which the original bundles initially remain discrete, most commonly in a tetrapolar arrangement, with four separate strands. Followed further in the basipetal direction, the strands eventually fuse partly or completely, usually to form a collateral or amphivasal axial bundle which participates in a new structural cycle. Quantitative variation between different taxa includes a simple pattern in Ludovia, in which only bipolar bundles are developed. More elaborate forms have multipolar bundles with more than four separate strands. A systematically useful observation is that stem vasculature in Cyclanthus, representing the subfamily Cyclanthoideae, does not differ significantly from that in subfamily Carludovicoideae although there are some distinctive structural features.     

DEVELOPMENT AND ORGANIZATION OF THE PRIMARY VASCULAR SYSTEM IN POPULUS DELTOIDES ACCORDING TO PHYLLOTAXY

An actively growing cottonwood bud was embedded in epon-araldite and serially sectioned at 2 μm. The sections were analyzed microscopically with the optical shuttle system of Zimmermann and Tomlinson, and all data were quantitatively recorded relative to the apex and to leaf plastochron index (LPI). Analysis of the sections revealed an acropetally developing procambial system organized according to a precise phyllotaxy. Six procambial strands could be recognized and followed long before the leaf primordia that they would enter were evident at the apex. Origin of these strands coincided with developmental events both in the parent trace and its primordium and in the antecedent leaf on the same orthostichy. Once a primordium and its trace attained a certain stage of development, trace bundles began to develop basipetally from the primordium base. These trace bundles appeared to be the earliest progenitors of wood formation in cottonwood. It was concluded that the concept of residual meristem and its corollary, the hypothesis that acropetally developing procambial strands determine the inception sties of new primordia, apply to the cottonwood apex.     

The Susceptibility of Monocotyledons to Agrobacterium tumefaciens

The susceptibility of 257 monocotyledon species belonging to 139 genera and 27 families, has been tested and the literature on this subject reviewed. In contrast with dicotyledons and gymnosperms, monocotyledons are much less susceptible to Agrobacterium tumefaciens: only 3 % of the species of monocotyledons tested were host plants, whereas 60 % of the dicotyledons and gymnosperms were susceptible. Only the closely related monocotyledon orders Liliales (6 families) and Arales (1 family) contained species that were susceptible. These results are consistent with a proposed taxonomic relationship between the Liliales and the dicotyledons. A correlation between host biochemical capabilities and susceptibility to crown gall is suggested.     

兰花蕉花部维管束系统的解剖学研究

兰花蕉花梗的维管束分散排列．子房基部的维管束排成两部分,外方为一轮大维管束环,中央为分散排列的小维管束区。前者的纸管束进入子房壁,后者进入子房的中轴,形成股座纸管束;及至延长都以后,股座维管束逐渐消失．子房壁上的维管束较易识别的有心皮背束、心皮背束伴束和隔膜束．三束心皮背束经延长部最终进入花柱和柱头．心皮背束指心皮背束务与其紧靠的大维管束,三枚心皮背束伴束最终分别进入三枚外轮雄蓝．三枚隔膜束中远轴面的两枚分别进入两校内轮雄蕊,而近轴面的一枚伴随着第六枚雄蓝的缺失最后进入唇瓣中央．子房壁其余的维管束进入延长部后,先向外分出一轮纸管束进入花幕,余下的中央部分排成一轮心形的线管来环．该环远轴面的维管束分为两半分别进入两枚侧生花瓣;近轴面即心形凹陷一侧初为两轮即外轮大的维管束与内轮小的维管束,后排成一轮并与近轴面的隔膜束一同进入唇瓣．兰花蕉的唇瓣既为花瓣成员,又含一枚缺失的雄蓝维管束,与姜目已报道的只来自退化雄蕊的竹芋科的兜状结构和美人蕉科、姜科、闭鞘姜科的唇瓣有明显区别．在旅人蕉科尚未有研究资料的情况下,作者根据已有资料,对姜目雄蕊维管束系统来源和结构进行比较,初步认为在姜目的系统演化上,兰花蕉科与芭蕉料更近．     

THE RELATIONSHIPS BETWEEN THE PRIMARY VASCULAR SYSTEM AND PHYLLOTACTIC PATTERNS OF ANAGALLIS ARVENSIS (PRIMULACEAE)

The various primary vascular systems of shoots of Anagallis arvensis L. (Primulaceae) can be distinguished in relation to the number of leaves (two, three, or four) at each node. In this study, shoot segments (single intemodes and the nodes above them) were examined. The arrangement of segments within shoots was also recorded. The vasculature forms a closed system with the number of sets of bundles usually equal to twice the number of leaves. Irregularities are found in the following features of the system: the number of bundles composing leaf half-traces; occurrence of anastomosing bundles; the number of intemodes through which bundles extend; levels of leaf attachment to the stem at the node; and distribution of parenchyma within the vascular cylinder, which determines the number of bundles in sets and the number of bundle sets. The irregularities occur with different frequencies for segments exhibiting different phyllotactic patterns. Comparison of these frequencies leads to the following conclusions: anastomosing bundles occur only in decussate or trimerous shoot segments, whereas sets of bundles united within intemodes and displaced leaves occur only in tetramerous or trimerous ones; decrease of the number of bundles per leaf and displacement of leaves at the nodal level are correlated; variation between segments exhibiting the same phyllotactic pattern is greatest for trimerous, less for tetramerous, and least for decussate segments; the vascular system of decussate shoot segments is more stable than that of the other systems; and trimerous segments seem to be intermediate between the other two segment types.     

HISTOLOGICAL STUDIES ON THE COLEOPTILE. II. COMPARATIVE VASCULAR ANATOMY OF COLEOPTILES OF AVENA AND TRITICUM

The vascular bundles in the uppermost 1-4 mm of the coleoptiles of 9 varieties of Avena sativa, and also of Avena fatua L., all terminate essentially vertically with a small “cap” of tracheary elements. In Triticum vulgare Vill., by contrast, they terminate with a horizontal or downward-pointing section. This brings the two bundles more or less together and may result in their complete fusion, usually with a short vascular extension. In both genera the bundles contain one or more series of apparently active, undifferentiated cells. In the mature embryos the bundles are entirely procambial in nature, but xylem differentiation begins rapidly upon germination and proceeds towards the tip, which is reached by the time the coleoptile is 1.5 mm long; thereafter it proceeds basipetally and it may continue at the base after elongation has ceased there. The differentiation of stomata also appears to proceed basipetally. It may be deduced that the coleoptile cannot form lignin while in the embryo but begins to do so upon germination. Parallels are brought out between auxin production first by the endosperm and then by the tip, on the one hand, and lignification in the xylem and in the stomata, on the other.     

COMPARATIVE MORPHOLOGY OF THE PRIMARY VASCULAR SYSTEMS IN SOME SPECIES OF ROSACEAE AND LEGUMINOSAE

The structural patterns of the primary vascular systems in some species of Leguminosae and Rosaceae have been determined by tracing the longitudinal course of the vascular bundles in terminal stem segments. These systems are interpreted as consisting of sympodia. Each sympodium is composed of an axial bundle which is continuous through the length of the segment and from which arise trace bundles that supply leaves and axillary buds. A compact arrangement of vascular bundles seems to correlate with the woody habit. Regardless of the degree of compactness of the primary vascular system, the structural identity of the individual sympodia is maintained. The total number of vascular bundles at a particular level is related to the number of axial bundles in the system, the number of traces per leaf and per axillary bud, and the number of internodes traversed by the traces prior to entering a lateral appendage. Shrubs and trees have more vascular bundles than herbs. Data from this study and the literature indicate that the vascular system is predominantly of the open type in dicotyledonous plants which have helically arranged leaves and, further, that in such plants with a 3-trace, trilacunar nodal structure, the number of sympodia coincides with the number of orthostichies (which is also the denominator of the phyllotactic fraction). In open systems leaf gaps cannot be morphologically delimited. Because of the resemblance of the open type of angiosperm vascular system to that of certain gymnosperms, previously interpreted to have evolved from a protostele, we suggest that the eustele of angiosperms is homologous with the stele of gymnosperms. We believe, also, that angiosperms, like gymnosperms, are probably not characterized by leaf gaps of filicinean type. We provide, furthermore, a rationale for the view that the axial bundle of a sympodium is a cauline structure.     

ONTOGENY OF THE PRIMARY THICKENING MERISTEM IN SEEDLINGS OF BOUGAINVILLEA SPECTABILIS

Anomalous secondary thickening occurs in the main axis of Bougainvillea spectabilis as a result of a primary thickening meristem which differentiates in pericycle. The primary thickening meristem first appears in the base of the primary root about 6 days after germination and differentiates acropetally as the root elongates. It begins differentiating from the base of the hypocotyl toward the shoot apex about 33 days after germination. The primary thickening meristem is first observable at the base of the first internode about 60 days after germination. It then becomes a cylinder in the main axis of the seedling. No stelar cambial cylinder forms in the primary root, hypocotyl, or stem because vascular cambium differentiation occurs neither in the pericycle opposite xylem points in the primary root nor in interfascicular parenchyma in the hypocotyl or stem. The primary vascular system of the stem appears anomalous because an inner and an outer ring of vascular bundles differentiate in the stele. Bundles of the inner ring anastomose in internodes, whereas those of the outer ring do not. Desmogen strands each of which is composed of phloem, xylem with both tracheids and vessels, and a desmogic cambium, differentiate from prodesmogen strands in conjunctive tissue. The parenchymatous cells surrounding desmogen strands then differentiate into elongated simple-pitted fibers and thick-walled fusiform cells that are about the same length as the primary thickening meristem initials.     

Potential allelochemicals from Sambucus nigra.

Twenty-four aromatic metabolites belonging to cyanogenins, lignans, flavonoids, and phenolic glycosides were obtained from Sambucus nigra. Structures were determined on the basis of their spectroscopic features. Two compounds have been isolated and identified as (2S)-2-O-beta-D-glucopyranosyl-2-hydroxyphenylacetic acid and benzyl 2-O-beta-D-glucopyranosyl-2,6-dihydroxybenzoate. All the compounds have been assayed on dicotyledons Lactuca sativa (lettuce) and Raphanus sativus (radish) and monocotyledon Allium cepa (onion) to test their stimulatory or inhibitory effects on seed germination and radicle elongation. Cyanogenins have a mainly inhibiting effect while lignans stimulate the growth. Some compounds show different effects on dicotyledons and monocotyledons.     

RE-EVALUATION OF CERTAIN KEY RELATIONSHIPS IN THE ALISMATIDAE: FLORAL ORGANOGENESIS OF SCHEUCHZERIA PALUSTRIS (SCHEUCHZERIACEAE)

Transition to flowering in the North-temperate bog plant Scheuchzeria palustris occurs in early May and results in the formation of a simple raceme with six flowers. Five of the flowers are subtended by large foliar bracts, while the sixth and last-formed flower on the inflorescence remains ebracteate. The individual flowers develop along a clearly trimerous pattern. The three outer tepals develop first, arising almost simultaneously at the periphery of the triangular floral apex. They are followed closely by the development of the three anti-tepalous outer stamens. The three inner tepals are next in the developmental sequence, alternating with the outer whorl of tepal-stamen pairs but arising at a slightly higher level on the floral meristem. Three inner stamens are initiated opposite the inner tepal primordia. Finally, three gynoecial primordia are initiated on the remaining central portion of the floral apex and alternating with the inner whorl of tepal-stamen pairs. Each carpel develops at first as a horseshoe-shaped structure. Two ovules form in each carpel, initiating on the adaxial margin of the carpel wall. Histogenesis of all floral appendages involves initially periclinal divisions in the second tunica layer followed by corresponding anticlinal divisions in the first tunica layer and concurrent activity in the underlying corpus. Separate procambial strands differentiate acropetally from the inflorescence axis to each tepal-stamen pair and then bifurcate. The vascular connection to the gynoecium develops directly from the strands in the tepal-stamen pairs. The results of this developmental study of the flower of S. palustris have a significant bearing on the positioning of this and related taxa within the Alismatidae and on the speculation of the phylogeny of the monocotyledon flower.     

金嘴蝎尾蕉花部维管束系统的解剖学研究

利用石蜡切片技术对蝎尾蕉科代表植物金嘴蝎尾蕉（Heliconia rostrata Ruiz＆Pavon）的花部维管束系统进行了解剖学研究。结果表明,心皮背束在延长部的基部分裂为内外2分支,内方分支与胎座维管束汇合后进入花柱,远轴面2枚外方分支在延长部的顶部分裂为2～4束进入远轴面2枚外轮雄蕊,而近轴面1枚外方分支则进入退化结构成为其中脉;隔膜束在延长部顶部亦分裂为3～5束,最终分别进入3枚内轮雄蕊;子房壁其它维管束最终进入花被片。本研究认为金嘴蝎尾蕉花部花瓣状退化结构与另外2枚外轮雄蕊具有完全相同的维管束系统来源,应属于雄蕊成员,且支持Kress关于蝎尾蕉科是姜群的姊妹群,区别于芭蕉群其它3科的观点。